Plant Transcription Factor Database
Previous version: v3.0
Transcription Factor Information
Basic Information | Signature Domain | Sequence | 
Basic Information? help Back to Top
TF ID AT1G21970.1
Common NameAtLEC1, EMB212, EMB 212, LEC1, NFYB9, NF-YB9, T26F17.20
Taxonomic ID
Taxonomic Lineage
cellular organisms; Eukaryota; Viridiplantae; Streptophyta; Streptophytina; Embryophyta; Tracheophyta; Euphyllophyta; Spermatophyta; Magnoliophyta; Mesangiospermae; eudicotyledons; Gunneridae; Pentapetalae; rosids; malvids; Brassicales; Brassicaceae; Camelineae; Arabidopsis
Family NF-YB
Protein Properties Length: 238aa    MW: 26070 Da    PI: 5.9417
Description NF-YB family protein
Gene Model
Gene Model ID Type Source Coding Sequence
AT1G21970.1genomeTAIRView CDS
Signature Domain? help Back to Top
Signature Domain
No. Domain Score E-value Start End HMM Start HMM End
        NF-YB   2 reqdrflPianvsrimkkvlPanakiskdaketvqecvsefisfvtseasdkcqrekrktingddllwalatlGfedyveplkvylkkyrelegek 97 
                  89*******************************************************************************************997 PP

Protein Features ? help Back to Top
3D Structure
Database Entry ID E-value Start End InterPro ID Description
PfamPF008081.8E-2663127IPR003958Transcription factor CBF/NF-Y/archaeal histone domain
PRINTSPR006153.0E-1691109No hitNo description
PROSITE patternPS00685094110IPR003956Transcription factor, NFYB/HAP3, conserved site
PRINTSPR006153.0E-16110128No hitNo description
PRINTSPR006153.0E-16129147No hitNo description
Gene Ontology ? help Back to Top
GO Term GO Category GO Description
GO:0009738Biological Processabscisic acid-activated signaling pathway
GO:0009785Biological Processblue light signaling pathway
GO:0010262Biological Processsomatic embryogenesis
GO:0045723Biological Processpositive regulation of fatty acid biosynthetic process
GO:0045893Biological Processpositive regulation of transcription, DNA-templated
GO:0005634Cellular Componentnucleus
GO:0003700Molecular Functiontranscription factor activity, sequence-specific DNA binding
GO:0005515Molecular Functionprotein binding
GO:0043565Molecular Functionsequence-specific DNA binding
GO:0046982Molecular Functionprotein heterodimerization activity
Plant Ontology ? help Back to Top
PO Term PO Category PO Description
PO:0000011anatomycultured somatic plant embryo
PO:0001078developmental stageplant embryo cotyledonary stage
PO:0001180developmental stageplant proembryo stage
PO:0001185developmental stageplant embryo globular stage
PO:0004507developmental stageplant embryo bilateral stage
Sequence ? help Back to Top
Protein Sequence    Length: 238 aa     Download sequence    Send to blast
3D Structure ? help Back to Top
PDB ID Evalue Query Start Query End Hit Start Hit End Description
Search in ModeBase
Expression -- UniGene ? help Back to Top
UniGene ID E-value Expressed in
Expression -- Microarray ? help Back to Top
Source ID E-value
Expression AtlasAT1G21970-
Expression -- Description ? help Back to Top
Source Description
UniprotDEVELOPMENTAL STAGE: Accumulates during seed development in embryo cell types and in endosperm tissue. {ECO:0000269|PubMed:9657152}.
UniprotTISSUE SPECIFICITY: Expressed in green siliques. Present in etiolated seedlings. {ECO:0000269|PubMed:11250072, ECO:0000269|PubMed:17322342}.
Functional Description ? help Back to Top
Source Description
TAIRTranscriptional activator of genes required for both embryo maturation and cellular differentiation. Sequence is similar to HAP3 subunit of the CCAAT-box binding factor. HAP3 subunit is divided into three domains: an amino-terminal A domain, a central B domain, and a carboxyl-terminal C domain. LEC1 shared high similarity with other HAP3 homologs only in central, B domain. LEC1 is required for the specification of cotyledon identity and the completion of embryo maturation. It was sufficient to induce embryogenic programs in vegetative cells, suggesting that LEC1 is a major embryonic regulator that mediates the switch between embryo and vegetative development. Mutants are desiccation intolerant, have trichomes on cotyledons and exhibit precocious meristem activation. Levels of the ABI3 and FUS3 transcripts were significantly reduced in developing siliques of the lec1-1 mutants, indicating that LEC1 down-regulates FUS3 and ABI3.When LEC1 is overexpressed from an inducible promoter, the expression of numerous genes involved in fatty acid biosynthesis is increased suggesting a role in positive regulation of FA biosynthesis.
UniProtComponent of the NF-Y/HAP transcription factor complex. The NF-Y complex stimulates the transcription of various genes by recognizing and binding to a CCAAT motif in promoters. Acts as a central regulator of the embryogenesis. Required for the speciation of cotyledon identity and the completion of embryo maturation. Controls seed storage protein genes through the regulation of FUS3 and ABI3. Involved in the blue light (BL) and abscisic acid (ABA) signaling pathways. {ECO:0000269|PubMed:12578989, ECO:0000269|PubMed:15695450, ECO:0000269|PubMed:17322342, ECO:0000269|PubMed:9657152}.
Function -- GeneRIF ? help Back to Top
  1. We suggest that LEC1 controls the expression of the SSP genes in a hierarchical manner, which involves ABI3 and FUS3.
    [PMID: 15695450]
  2. Cultures of the lec mutants formed somatic embryos at a low frequency. Moreover, somatic embryos were formed from callus tissue through an indirect route.
    [PMID: 16034595]
  3. A study of the network of transcription factors that regulates gene expression in A. thaliana is described and discussed.
    [PMID: 17158584]
  4. LEC1 positively regulates fatty acid biosynthesis genes. Genes involved in glycolysis and lipid accumulation are also up-regulated.
    [PMID: 18689444]
  5. LEC1 and LEC1-LIKE activate the CRC and SUCROSE SYNTHASE 2 (SUS2) promoters in combination with an NF-YC subunit through the interaction with a seed-specific ABA-response element (ABRE) binding bZIP factor, bZIP67.
    [PMID: 19207209]
  6. LEC1 appears to be an integrator of various regulatory events, involving the transcription factor itself as well as light and hormone signalling, especially during somatic and early zygotic embryogenesis.
    [PMID: 22429691]
  7. Data confirms LEC1's known participation in the regulation of somatic embryogenesis, but also indicates additional roles in embryonic and extra-embryonic cell elongation
    [PMID: 23073004]
  8. NUCLEAR FACTOR Y transcription factors have both opposing and additive roles in abscisic acid-mediated seed germination.
    [PMID: 23527203]
  9. NUCLEAR FACTOR Y (NF-Y) binds a CCAAT box in the distal enhancer element and that CCAAT disruption dramatically reduces FT promoter activity
    [PMID: 24610724]
  10. In addition to VAL B3 factors, epigenetic mechanisms are implicated in maintaining repression of LEC1/AFL (LAFL)network during vegetative development
    [PMID: 24902838]
Cis-element ? help Back to Top
Regulation -- PlantRegMap ? help Back to Top
Source Upstream Regulator Target Gene
Regulation -- ATRM (Manually Curated Upstream Regulators) ? help Back to Top
Source Upstream Regulator (A: Activate/R: Repress)
ATRM AT1G54060 (R), AT2G17950 (R), AT3G27785 (A), AT5G40360 (A)
Regulation -- ATRM (Manually Curated Target Genes) ? help Back to Top
Source Target Gene (A: Activate/R: Repress)
ATRM AT1G06180(R), AT1G28300(A), AT1G62290(A), AT2G17090(A), AT2G36270(A), AT2G41260(A), AT3G24650(A), AT3G26790(A), AT3G51810(R), AT3G54320(A), AT4G27160(A), AT5G66400(A)
Regulation -- Hormone ? help Back to Top
Source Hormone
AHDabscisic acid
Interaction ? help Back to Top
Source Intact With
BioGRIDAT1G54830, AT1G56170
IntActSearch Q9SFD8
Phenotype -- Disruption Phenotype ? help Back to Top
Source Description
UniProtDISRUPTION PHENOTYPE: Altered response to blue light (BL) and abscisic acid (ABA). {ECO:0000269|PubMed:17322342}.
Phenotype -- Mutation ? help Back to Top
Source ID
T-DNA ExpressAT1G21970
Annotation -- Nucleotide ? help Back to Top
Source Hit ID E-value Description
GenBankAB4934740.0AB493474.1 Arabidopsis thaliana At1g21970 mRNA for hypothetical protein, partial cds, clone: RAAt1g21970.
Annotation -- Protein ? help Back to Top
Source Hit ID E-value Description
RefseqNP_173616.21e-179nuclear transcription factor Y subunit B-9
SwissprotQ9SFD80.0NFYB9_ARATH; Nuclear transcription factor Y subunit B-9
TrEMBLD7MX191e-158D7MX19_ARALL; Putative uncharacterized protein
STRINGAT1G21970.11e-178(Arabidopsis thaliana)
Orthologous Group ? help Back to Top
LineageOrthologous Group IDTaxa NumberGene Number
Representative plantOGRP16817170
Publications ? help Back to Top
  1. Wobus U,Weber H
    Seed maturation: genetic programmes and control signals.
    Curr. Opin. Plant Biol., 1999. 2(1): p. 33-8
  2. Ogas J,Kaufmann S,Henderson J,Somerville C
    PICKLE is a CHD3 chromatin-remodeling factor that regulates the transition from embryonic to vegetative development in Arabidopsis.
    Proc. Natl. Acad. Sci. U.S.A., 1999. 96(24): p. 13839-44
  3. Rohde A, et al.
    ABI3 affects plastid differentiation in dark-grown Arabidopsis seedlings.
    Plant Cell, 2000. 12(1): p. 35-52
  4. Kurup S,Jones HD,Holdsworth MJ
    Interactions of the developmental regulator ABI3 with proteins identified from developing Arabidopsis seeds.
    Plant J., 2000. 21(2): p. 143-55
  5. Wehmeyer N,Vierling E
    The expression of small heat shock proteins in seeds responds to discrete developmental signals and suggests a general protective role in desiccation tolerance.
    Plant Physiol., 2000. 122(4): p. 1099-108
  6. Tsukaya H,Shoda K,Kim GT,Uchimiya H
    Heteroblasty in Arabidopsis thaliana (L.) Heynh.
    Planta, 2000. 210(4): p. 536-42
  7. Vicient CM,Bies-Etheve N,Delseny M
    Changes in gene expression in the leafy cotyledon1 (lec1) and fusca3 (fus3) mutants of Arabidopsis thaliana L.
    J. Exp. Bot., 2000. 51(347): p. 995-1003
  8. Riechmann JL, et al.
    Arabidopsis transcription factors: genome-wide comparative analysis among eukaryotes.
    Science, 2000. 290(5499): p. 2105-10
  9. Gusmaroli G,Tonelli C,Mantovani R
    Regulation of the CCAAT-Binding NF-Y subunits in Arabidopsis thaliana.
    Gene, 2001. 264(2): p. 173-85
  10. Hong SW,Vierling E
    Hsp101 is necessary for heat tolerance but dispensable for development and germination in the absence of stress.
    Plant J., 2001. 27(1): p. 25-35
  11. Gusmaroli G,Tonelli C,Mantovani R
    Regulation of novel members of the Arabidopsis thaliana CCAAT-binding nuclear factor Y subunits.
    Gene, 2002. 283(1-2): p. 41-8
  12. Zuo J,Niu QW,Frugis G,Chua NH
    The WUSCHEL gene promotes vegetative-to-embryonic transition in Arabidopsis.
    Plant J., 2002. 30(3): p. 349-59
  13. Zhang S,Wong L,Meng L,Lemaux PG
    Similarity of expression patterns of knotted1 and ZmLEC1 during somatic and zygotic embryogenesis in maize ( Zea mays L.).
    Planta, 2002. 215(2): p. 191-4
  14. Finkelstein RR,Gampala SS,Rock CD
    Abscisic acid signaling in seeds and seedlings.
    Plant Cell, 2002. 14 Suppl: p. S15-45
  15. West M, et al.
    LEAFY COTYLEDON1 Is an Essential Regulator of Late Embryogenesis and Cotyledon Identity in Arabidopsis.
    Plant Cell, 1994. 6(12): p. 1731-1745
  16. Meinke DW,Franzmann LH,Nickle TC,Yeung EC
    Leafy Cotyledon Mutants of Arabidopsis.
    Plant Cell, 1994. 6(8): p. 1049-1064
  17. Kwong RW, et al.
    LEAFY COTYLEDON1-LIKE defines a class of regulators essential for embryo development.
    Plant Cell, 2003. 15(1): p. 5-18
  18. Brocard-Gifford IM,Lynch TJ,Finkelstein RR
    Regulatory networks in seeds integrating developmental, abscisic acid, sugar, and light signaling.
    Plant Physiol., 2003. 131(1): p. 78-92
  19. Lee H,Fischer RL,Goldberg RB,Harada JJ
    Arabidopsis LEAFY COTYLEDON1 represents a functionally specialized subunit of the CCAAT binding transcription factor.
    Proc. Natl. Acad. Sci. U.S.A., 2003. 100(4): p. 2152-6
  20. Dean Rider S, et al.
    Coordinate repression of regulators of embryonic identity by PICKLE during germination in Arabidopsis.
    Plant J., 2003. 35(1): p. 33-43
  21. Baumbusch LO,Hughes DW,Galau GA,Jakobsen KS
    LEC1, FUS3, ABI3 and Em expression reveals no correlation with dormancy in Arabidopsis.
    J. Exp. Bot., 2004. 55(394): p. 77-87
  22. Castelli V, et al.
    Whole genome sequence comparisons and "full-length" cDNA sequences: a combined approach to evaluate and improve Arabidopsis genome annotation.
    Genome Res., 2004. 14(3): p. 406-13
  23. Yazawa K,Takahata K,Kamada H
    Isolation of the gene encoding Carrot leafy cotyledon1 and expression analysis during somatic and zygotic embryogenesis.
    Plant Physiol. Biochem., 2004. 42(3): p. 215-23
  24. Kagaya Y, et al.
    LEAFY COTYLEDON1 controls seed storage protein genes through its regulation of FUSCA3 and ABSCISIC ACID INSENSITIVE3.
    Plant Cell Physiol., 2005. 46(3): p. 399-406
  25. Gaj MD,Zhang S,Harada JJ,Lemaux PG
    Leafy cotyledon genes are essential for induction of somatic embryogenesis of Arabidopsis.
    Planta, 2005. 222(6): p. 977-88
  26. Santos Mendoza M,Dubreucq B,Miquel M,Caboche M,Lepiniec L
    LEAFY COTYLEDON 2 activation is sufficient to trigger the accumulation of oil and seed specific mRNAs in Arabidopsis leaves.
    FEBS Lett., 2005. 579(21): p. 4666-70
  27. Fambrini M, et al.
    Characterization of LEAFY COTYLEDON1-LIKE gene in Helianthus annuus and its relationship with zygotic and somatic embryogenesis.
    Dev. Genes Evol., 2006. 216(5): p. 253-64
  28. Shen B,Sinkevicius KW,Selinger DA,Tarczynski MC
    The homeobox gene GLABRA2 affects seed oil content in Arabidopsis.
    Plant Mol. Biol., 2006. 60(3): p. 377-87
  29. To A, et al.
    A network of local and redundant gene regulation governs Arabidopsis seed maturation.
    Plant Cell, 2006. 18(7): p. 1642-51
  30. Casson SA,Lindsey K
    The turnip mutant of Arabidopsis reveals that LEAFY COTYLEDON1 expression mediates the effects of auxin and sugars to promote embryonic cell identity.
    Plant Physiol., 2006. 142(2): p. 526-41
  31. Wenkel S, et al.
    CONSTANS and the CCAAT box binding complex share a functionally important domain and interact to regulate flowering of Arabidopsis.
    Plant Cell, 2006. 18(11): p. 2971-84
  32. Suzuki M,Wang HH,McCarty DR
    Repression of the LEAFY COTYLEDON 1/B3 regulatory network in plant embryo development by VP1/ABSCISIC ACID INSENSITIVE 3-LIKE B3 genes.
    Plant Physiol., 2007. 143(2): p. 902-11
  33. Cao X, et al.
    Abscisic acid and stress signals induce Viviparous1 expression in seed and vegetative tissues of maize.
    Plant Physiol., 2007. 143(2): p. 720-31
  34. Warpeha KM, et al.
    The GCR1, GPA1, PRN1, NF-Y signal chain mediates both blue light and abscisic acid responses in Arabidopsis.
    Plant Physiol., 2007. 143(4): p. 1590-600
  35. Wang H,Guo J,Lambert KN,Lin Y
    Developmental control of Arabidopsis seed oil biosynthesis.
    Planta, 2007. 226(3): p. 773-83
  36. Tanaka M,Kikuchi A,Kamada H
    The Arabidopsis histone deacetylases HDA6 and HDA19 contribute to the repression of embryonic properties after germination.
    Plant Physiol., 2008. 146(1): p. 149-61
  37. Yazawa K,Kamada H
    Identification and characterization of carrot HAP factors that form a complex with the embryo-specific transcription factor C-LEC1.
    J. Exp. Bot., 2007. 58(13): p. 3819-28
  38. Alemanno L, et al.
    Characterization of leafy cotyledon1-like during embryogenesis in Theobroma cacao L.
    Planta, 2008. 227(4): p. 853-66
  39. Suzuki M, et al.
    The Maize Viviparous8 locus, encoding a putative ALTERED MERISTEM PROGRAM1-like peptidase, regulates abscisic acid accumulation and coordinates embryo and endosperm development.
    Plant Physiol., 2008. 146(3): p. 1193-206
  40. Stone SL, et al.
    Arabidopsis LEAFY COTYLEDON2 induces maturation traits and auxin activity: Implications for somatic embryogenesis.
    Proc. Natl. Acad. Sci. U.S.A., 2008. 105(8): p. 3151-6
  41. Santos-Mendoza M, et al.
    Deciphering gene regulatory networks that control seed development and maturation in Arabidopsis.
    Plant J., 2008. 54(4): p. 608-20
  42. Mu J, et al.
    LEAFY COTYLEDON1 is a key regulator of fatty acid biosynthesis in Arabidopsis.
    Plant Physiol., 2008. 148(2): p. 1042-54
  43. Wang X, et al.
    Overexpression of PGA37/MYB118 and MYB115 promotes vegetative-to-embryonic transition in Arabidopsis.
    Cell Res., 2009. 19(2): p. 224-35
  44. Gao MJ, et al.
    Repression of seed maturation genes by a trihelix transcriptional repressor in Arabidopsis seedlings.
    Plant Cell, 2009. 21(1): p. 54-71
  45. Yamamoto A, et al.
    Arabidopsis NF-YB subunits LEC1 and LEC1-LIKE activate transcription by interacting with seed-specific ABRE-binding factors.
    Plant J., 2009. 58(5): p. 843-56
  46. Huang Y, et al.
    Probing the endosperm gene expression landscape in Brassica napus.
    BMC Genomics, 2009. 10: p. 256
  47. Sugliani M,Rajjou L,Clerkx EJ,Koornneef M,Soppe WJ
    Natural modifiers of seed longevity in the Arabidopsis mutants abscisic acid insensitive3-5 (abi3-5) and leafy cotyledon1-3 (lec1-3).
    New Phytol., 2009. 184(4): p. 898-908
  48. Le BH, et al.
    Global analysis of gene activity during Arabidopsis seed development and identification of seed-specific transcription factors.
    Proc. Natl. Acad. Sci. U.S.A., 2010. 107(18): p. 8063-70
  49. Mitsuda N, et al.
    Efficient yeast one-/two-hybrid screening using a library composed only of transcription factors in Arabidopsis thaliana.
    Plant Cell Physiol., 2010. 51(12): p. 2145-51
  50. Yang Y,Karlson DT
    Overexpression of AtCSP4 affects late stages of embryo development in Arabidopsis.
    J. Exp. Bot., 2011. 62(6): p. 2079-91
  51. Tan H, et al.
    Enhanced seed oil production in canola by conditional expression of Brassica napus LEAFY COTYLEDON1 and LEC1-LIKE in developing seeds.
    Plant Physiol., 2011. 156(3): p. 1577-88
  52. Tang X, et al.
    Synergistic repression of the embryonic programme by SET DOMAIN GROUP 8 and EMBRYONIC FLOWER 2 in Arabidopsis seedlings.
    J. Exp. Bot., 2012. 63(3): p. 1391-404
  53. Hackenberg D, et al.
    Studies on differential nuclear translocation mechanism and assembly of the three subunits of the Arabidopsis thaliana transcription factor NF-Y.
    Mol Plant, 2012. 5(4): p. 876-88
  54. Junker A, et al.
    Elongation-related functions of LEAFY COTYLEDON1 during the development of Arabidopsis thaliana.
    Plant J., 2012. 71(3): p. 427-42
  55. Zhang H,Bishop B,Ringenberg W,Muir WM,Ogas J
    The CHD3 remodeler PICKLE associates with genes enriched for trimethylation of histone H3 lysine 27.
    Plant Physiol., 2012. 159(1): p. 418-32
  56. Calvenzani V, et al.
    Interactions and CCAAT-binding of Arabidopsis thaliana NF-Y subunits.
    PLoS ONE, 2012. 7(8): p. e42902
  57. Mu J,Tan H,Hong S,Liang Y,Zuo J
    Arabidopsis transcription factor genes NF-YA1, 5, 6, and 9 play redundant roles in male gametogenesis, embryogenesis, and seed development.
    Mol Plant, 2013. 6(1): p. 188-201
  58. Meinke DW
    A survey of dominant mutations in Arabidopsis thaliana.
    Trends Plant Sci., 2013. 18(2): p. 84-91
  59. Junker A,B
    Multifunctionality of the LEC1 transcription factor during plant development.
    Plant Signal Behav, 2012. 7(12): p. 1718-20
  60. Zhang JJ,Xue HW
    OsLEC1/OsHAP3E participates in the determination of meristem identity in both vegetative and reproductive developments of rice.
    J Integr Plant Biol, 2013. 55(3): p. 232-49
  61. Wang F,Perry SE
    Identification of direct targets of FUSCA3, a key regulator of Arabidopsis seed development.
    Plant Physiol., 2013. 161(3): p. 1251-64
  62. Li-Beisson Y, et al.
    Acyl-lipid metabolism.
    Arabidopsis Book, 2013. 11: p. e0161
  63. Kumimoto RW, et al.
    NUCLEAR FACTOR Y transcription factors have both opposing and additive roles in ABA-mediated seed germination.
    PLoS ONE, 2013. 8(3): p. e59481
  64. Kirkbride RC,Fischer RL,Harada JJ
    LEAFY COTYLEDON1, a key regulator of seed development, is expressed in vegetative and sexual propagules of Selaginella moellendorffii.
    PLoS ONE, 2013. 8(6): p. e67971
  65. Cao S, et al.
    A distal CCAAT/NUCLEAR FACTOR Y complex promotes chromatin looping at the FLOWERING LOCUS T promoter and regulates the timing of flowering in Arabidopsis.
    Plant Cell, 2014. 26(3): p. 1009-17
  66. Jia H,Suzuki M,McCarty DR
    Regulation of the seed to seedling developmental phase transition by the LAFL and VAL transcription factor networks.
    Wiley Interdiscip Rev Dev Biol, 2014 Jan-Feb. 3(1): p. 135-45
  67. Jin J, et al.
    An Arabidopsis Transcriptional Regulatory Map Reveals Distinct Functional and Evolutionary Features of Novel Transcription Factors.
    Mol. Biol. Evol., 2015. 32(7): p. 1767-73
  68. Perry SE,Nichols KW,Fernandez DE
    The MADS domain protein AGL15 localizes to the nucleus during early stages of seed development.
    Plant Cell, 1996. 8(11): p. 1977-89
  69. Parcy F,Valon C,Kohara A,Miséra S,Giraudat J
    The ABSCISIC ACID-INSENSITIVE3, FUSCA3, and LEAFY COTYLEDON1 loci act in concert to control multiple aspects of Arabidopsis seed development.
    Plant Cell, 1997. 9(8): p. 1265-77
  70. Lotan T, et al.
    Arabidopsis LEAFY COTYLEDON1 is sufficient to induce embryo development in vegetative cells.
    Cell, 1998. 93(7): p. 1195-205
  71. Kirik V,K
    Two novel MYB homologues with changed expression in late embryogenesis-defective Arabidopsis mutants.
    Plant Mol. Biol., 1998. 37(5): p. 819-27
  72. Kirik V,K
    Ectopic expression of a novel MYB gene modifies the architecture of the Arabidopsis inflorescence.
    Plant J., 1998. 13(6): p. 729-42
  73. Wolkers WF,Alberda M,Koornneef M,L
    Properties of proteins and the glassy matrix in maturation-defective mutant seeds of Arabidopsis thaliana.
    Plant J., 1998. 16(2): p. 133-43